A New Species of Marmara (Lepidoptera: Gracillariidae: Marmarinae), with an Annotated List of Known Hostplants for the Genus

Zootaxa 4337 (2): 198–222 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4337.2.2 http://zoobank.org/urn:lsid:zoobank.org:pub:4DC4B753-3A04-41F0-94D7-C25FB9F4A713 A new species of Marmara (Lepidoptera: Gracillariidae: Marmarinae), with an Annotated List of Known Hostplants for the Genus

CHARLES S. EISEMAN1, DONALD R. DAVIS2, JULIA A. BLYTH1, DAVID L. WAGNER3, MICHAEL W. PALMER4 & TRACY S. FELDMAN5 1276 Old Wendell Rd., Northfield, MA 01360-9674, U.S.A. E-mail: [email protected] 2Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012 MRC 105, Washington, D.C. 20013-7012, U.S.A. Email: [email protected] 3Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269-3043, U.S.A. Email: [email protected] 4Department of Plant Biology, Ecology and Evolution, 301 Physical Sciences, Oklahoma State University, Stillwater, OK 74078-3013, U.S.A. E-mail: [email protected] 5 Department of Natural and Life Sciences, St. Andrews University, a branch of Webber International University, 1700 Dogwood Mile, Laurinburg, NC 28352-5521, U.S.A. E-mail: [email protected]

Abstract

Larvae of the New World gracillariid moth genus Marmara are primarily stem/bark miners, with some species mining in leaves or fruits. We describe a new species, M. viburnella Eiseman & Davis, which feeds on Viburnum, initially mining the leaves but completing development as a stem miner. The type series is from Nantucket Island, Massachusetts, with observations of leaf mines indicating the species is widespread in the eastern USA. Combining previously published data, our own observations, and other sources, we present a list of known Marmara hostplants, many of which represent undescribed species.

Key words: Ageniaspis, Quadrastichus, barkminer, leafminer, stem miner, Viburnum

Introduction

The genus Marmara Clemens (Lepidoptera: Gracillariidae: Marmarinae) includes 19 described North American species and five from South America. Larvae of most species are stem miners, typically in bark of woody plants, with a few mining instead (or additionally) in leaves or fruits. When mature, larvae of all the leaf and fruit feeders, and most of the stem feeders, molt to a non-feeding final instar that cuts a transverse semicircular slit at the end of the mine and wanders for some distance before spinning a cocoon. This cocoon may be spun on a leaf, in a bark crevice, or in leaf litter, and is adorned with numerous pearly globules or “bubbles” extruded from the anus (Wagner et al. 2000). A few stem-mining species instead cut a much larger, lateral, semicircular flap at the end of the mine and spin an unadorned cocoon on the underside of this flap, causing the overlying stem tissues to buckle over the cocoon (Braun 1915; Vinal 1917; De Gryse 1943; Fitzgerald 1973; Wagner et al. 2000). One Marmara species from the southwestern USA and all of the South American species were described by E. Meyrick without knowledge of larval biology, although adults reared from cacao pods were subsequently determined as M. isortha (Meyrick) (Bondar 1939). Marmara gulosa Guillén & Davis is polyphagous, mining in peels, shoots, and occasionally leaves in California and Arizona (Guillén et al. 2001; Neff 2002; Semet 2010). Known hosts include an array of cultivated plants as well as some native species, although there has yet to be a focused effort to assess its host range on natives. As far as is known, the remaining described species each feed on plants of one genus or two closely related genera. Marmara smilacisella (Chambers) mines leaves of Smilax L. (Smilacaceae), M. arbutiella Busck mines leaves and occasionally green stems of Arbutus L. (Ericaceae), and the rest are apparently exclusively stem miners (Wagner et al. 2000; Guillén et al. 2001). A table of the host and tissue specificity of described Marmara species was presented by Guillén et al. (2001), and one additional species has been described since then (Davis et al. 2011).

198 Accepted by J. De Prins: 11 Jul. 2017; published: 18 Jul. 2017 Here we describe a new species that is intermediate in habits, with young larvae mining in leaves and later instars feeding in stems. The cocoon is spun under a bark flap cut at the mine terminus but is sometimes adorned with pearly bubbles. In the hope of stimulating further investigation of this genus, we present an annotated list of known and suspected Marmara host plants assembled from the literature and our own observations (Table 1).

Materials and methods

In June 2016, CSE and JAB spent several hours on Nantucket Island (Massachusetts, USA) searching Viburnum dentatum L. (Adoxaceae) plants for Marmara bark flaps, either gently peeling them off or using a knife to cut off stem portions that included the flaps. These were collected in plastic vials, which were checked daily for emerging adults. Adult moths were pinned, spread, and double mounted by JAB. After CSE described the external characters, the specimens were sent to DRD for dissection and illustrations, and were deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM). Parasitoids were preserved in 95% ethanol. Encyrtids were examined by R. L. Zuparko, Department of Entomology, California Academy of Sciences, San Francisco, and deposited in the Essig Museum of Entomology, University of California, Berkeley. Eulophids were examined by C. Hansson, Department of Zoology, Lund University, Sweden, and deposited in the Natural History Museum, London. We have followed Stevens (2016) for higher taxonomy of plants. Otherwise, plant taxonomy follows USDA, NRCS (2016), and literature records have been brought into agreement with this classification without comment except in ambiguous cases.

Marmara viburnella Eiseman & Davis, sp. nov. Figs. 1–11

Adult (Figs. 1, 10, 11). Wingspan ~6 mm. Head: Vestiture smooth; frons and vertex silvery white; back of head blackish. Eyes red in live specimens. Maxillary palpus white with prominent black tip; haustellum white; labial palpus with first two segments black (one specimen with inner surfaces white) and third segment white with a black ventral spot. Antenna dusky, paler beneath; pedicel with black scales (conspicuously elongated in female). Thorax: Shining blackish above, pale golden beneath. Forewing shining blackish with silvery markings: a broad transverse fascia in the basal ¼, gradually broadening dorsally; at ½, opposing dorsal and costal spots, separate (1 specimen) or narrowly joined in the middle (2 specimens); at ¾, a prominent costal spot (extending to middle of wing) and a smaller, opposing dorsal spot; a fainter costal spot of similar size at apex, mottled with blackish scales; fringe white. Hindwing dusky, slightly paler than forewing. Coxae white with black scales distally. Fore femur mostly black, with variable amounts of white proximally, laterally, and posteriorly; fore tibia mostly black, with variable amounts of white proximally, distally, and posteriorly; fore tarsus white with black bands anteriorly. Middle femur mostly black (including spurs); middle tibia black with a broad, white central band, more or less interrupted centrally with black scales; middle tarsus white with a few black scales. Hind femur white with a broad black band distally; hind tibia black with two broad white bands, contiguous with the white proximal and distal spurs; hind tarsus white with a broad black band proximally and a few black scales distally. Abdomen: Shining blackish dorsally; ventrally silvery with intersegmental boundaries black; anal tuft silvery (concolorous with abdomen in female). Male genitalia (Figs. 2A–B): Uncus absent. Tegumen a slender, dorsal arched band. Vinculum a moderately broad ventral band with anterior margin slightly curved caudally; anterior margin reflexed medially to form slender triangular lobe. Gnathos membranous and poorly defined. Valva separated nearly from base into three distinct lobes: a relatively short, slender, costal lobe bearing a dense comb of 18–20 short, stout spines; an elongate, slender, more lateral cucullar lobe that expands abruptly to form a setose, triangular distal lobe; and the largest, most ventral, valvular lobe that gradually broadens apically to a nearly truncate, inwardly curved apex. Phallus short, acute, with greatly inflated phallobase, approximately equal in length to distal, tubular portion of phallus.

FIGURE 2. Marmara viburnella male genitalia. 2a, Genital capsule with valvae, ventral view. 2b, Phallus, lateral view.

Female genitalia: Not examined. The abdomen is now missing from the single female specimen. Larva. Early instar as in Fig. 3; immature stages otherwise not examined. Cocoon. An oblong envelope of white silk, approximately 5–6 mm long and 2–3 mm wide, spun on the underside of a semicircular bark flap cut by the larva at the end of the mine; unadorned or with a cluster of 1–12 or so pearly bubbles near each end (Figs. 4–6). Type material. Holotype: ♂, UNITED STATES: Massachusetts: Nantucket Co.: Nantucket State Forest, 11.vi.2016, em. 2.vii.2016, C. S. Eiseman & J. A. Blyth, ex Viburnum dentatum, #CSE2692, slide USNM 34733, digital image captured (USNM 01325414). Paratypes: Same collection data as holotype, 1 ♂, em. 24.vi.2016, #CSE2625 (USNM); Nantucket, Lost Farm, 1 ♀, 12.vi.2016, em. 27.vi.2016, C. Eiseman, ex Viburnum dentatum, #CSE2642 (USNM).

200 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. FIGURES 3–11. Marmara viburnella. 3, early instar larva; 4, bark flap cut by the larva, under which the cocoon is spun; 5, cocoon of male paratype, with pupal exuviae protruding from right end; 6, cocoon with numerous pearly bubbles, found in Illinois; 7, the first documented leaf mine, from Tuckernuck Island, September 2011; 8, mine tract departing the leaf blade, visible as a brown line in the petiole and twig; 9, bark mine; 10, holotype male; 11, paratype male.

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 201 Distribution. Eastern United States and southeastern Canada. Adults have only been reared from cocoons collected on Nantucket Island, Massachusetts, but we have observed larval mines in mainland Massachusetts (Berkshire, Bristol, and Plymouth Counties), Connecticut (Hartford), Illinois (near Effingham), Louisiana (4 mi NE of Slidell), Maryland (Baltimore; Wilson 2014), North Carolina (Durham), Rhode Island (Block Island), Vermont (Mt. Mansfield), and Quebec (Gatineau Park). Etymology. The specific name is derived from the genus of the host plant, Viburnum L. Diagnosis. The forewing pattern is similar to that of several described Marmara species possessing darkly banded forewings, but M. viburnella is clearly distinct when larval biology is taken into account. No other species is known both to mine leaves and to pupate under a bark flap cut by the larva at the end of the mine. Further, each of the six species that is reported to pupate under a bark flap, in addition to using hosts belonging to other plant orders, has an adult with distinctly different forewing coloration: the wings of M. auratella Braun are “bronzy brown, with an almost golden luster under brilliant illumination” (Braun 1915); those of M. elotella (Busck) and the three ash feeders are predominantly white (Busck 1909; Fitzgerald 1973); and those of M. fasciella (Chambers) are banded with approximately equal parts white and pale brown (Chambers 1875). The male genitalia of M. viburnella are most similar to that of M. fulgidella (Clemens) in possessing a greatly inflated phallobase and inwardly curved apex of the valvular lobes. The forewing pattern of M. fulgidella is distinct in possessing much broader white fascia. Host plants. Adults have been reared from Viburnum dentatum L. (Adoxaceae). We have found leaf mines of the new species on V. lantanoides Michx., V. nudum L. var. cassinoides (L.) Torr. & A. Gray, and V. rafinesquianum Schult. Biology. In Massachusetts, adults emerge in June and early July and deposit eggs on the upper surfaces of leaves of Viburnum, one egg per leaf, typically over a prominent vein. The sap-feeding larvae hatch in early July and form meandering linear leaf mines, less than 1 mm wide (Figs. 7–8). The mine is at first epidermal and may appear whitish or like a tiny, shining snail trail, difficult to discern. Before long the mine deepens and becomes pale brown in color, with a very fine, central frass line visible under magnification. Eventually the mine enters the leaf midrib, either directly or by a side vein, and proceeds down the petiole and into the stem. We have found apparently occupied leaf mines in Massachusetts as late as 8 August, and have found completed leaf mines, with larvae already feeding in stems, as early as 19 July in Massachusetts and 30 June in Illinois. Once in the stem, the larva quickly tunnels deeper in the bark, and the mine (Fig. 9) usually is not externally visible for more than a few cm. Based on the known life histories of other bark-mining Marmara species, we presume that the larvae overwinter partially grown and finish feeding in the spring. The depth of the bark mine appears to vary throughout development; we have found fragmentary epidermal and deeper bark mines of various sizes, but much of the feeding evidently takes place in the cortex and is not visible on the surface. The mature larva cuts a semicircular flap in the bark (Fig. 4) and spins its cocoon on the underside of this (Fig. 6). Although we have found mines extending to within a few cm of the base of the stem, the pupation site tends to be well over 1 m aboveground, on a branch or a portion of the main stem that is less than 1 cm thick and has relatively smooth bark. Upon emergence of the adult (Figs. 10–11), the pupa is thrust through the cocoon near one end (Fig. 5). Parasitoids. Four Quadrastichus Girault adults (Hymenoptera: Eulophidae: Tetrastichinae) emerged from one cocoon. No keys exist for the identification of North American species in this genus, and there are likely numerous undescribed species in addition to the ten known from this continent (C. Hansson, in litt.). Six Ageniaspis Dahlbom adults (Encyrtidae: Encyrtinae) emerged from another cocoon, and another 23 emerged from two or more cocoons that were not isolated in separate rearing vials. They do not match any of the species recorded from northeastern North America (R. Zuparko, in litt.). Other Associates. We found several different arthropods hiding under bark flaps created by larvae of Marmara viburnella. These included two elongate-bodied springtails (Entomobryomorpha), Anurophorus cf. septentrionalis Palissa (Isotomidae) and Entomobrya nivalis (L.) (Entomobryidae), as well as two beetles (Coleoptera), Contacyphon Des Gozis sp. (Scirtidae) and Neapion Alonso-Zarazaga sp. (Brentidae). Remarks. The successful rearing of Marmara viburnella followed five years of investigation and failed attempts, during which the larval biology was gradually pieced together. The first known leaf mine on Viburnum dentatum was found on Tuckernuck Island by CSE on 10 September 2011 during a survey of leaf-mining and gall- forming insects of Nantucket County, Massachusetts, USA. Additional mines were found on Nantucket Island in August 2012, and since they were observed to proceed down the petioles and into the twigs, several twigs with

202 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. mined leaves were collected in resealable plastic bags. A single larva appeared in one of the bags, and photographs were shown to DRD and DLW, who agreed that it appeared to be an early instar Marmara. In June 2013, additional searching of V. dentatum by CSE and JAB revealed a few old bark mines, but no bark flaps were found and it was presumed that this species exits the mine to pupate after overwintering in the stem or roots. In late July 2014, V. dentatum plants with leaf mines were marked with pink flagging, and in December, five of these were dug, potted, and kept in an unheated shed over the winter, inside large sleeves of fine-meshed cloth. On 3 March 2015, the plants were brought indoors, and they were checked daily until June for emerging adults. No moths emerged, but on 30 June, CSE and JAB discovered leaf and stem mines with two associated bark flaps on V. dentatum west of Effingham, Illinois. Thus, in June 2016 CSE and JAB were newly motivated to search for bark flaps on Nantucket Island, and succeeded in collecting approximately 30 that appeared fresh (rejecting numerous others that were from previous years) scattered over five different sites. The leaf mine of Marmara viburnella is easily distinguished from other mines occurring on Viburnum, as no other insect produces a mine that proceeds down the petiole and into the stem. The only other known linear mine on Viburnum is one found by CSE and JAB on V. edul e (Michx.) Raf. in Washington. This mine was tightly coiled at the beginning, arcing back and forth in an area bounded by two lateral veins; eventually it crossed a vein and stretched out into a typical linear mine. The larva exited through a slit in the upper epidermis at the end of the mine. Frass was deposited in beaded strips along the sides (not in a thin central line as in M. viburnella), an arrangement characteristic of Agromyzidae (Diptera). We presume this mine to be the work of Liriomyza charada Lonsdale, which has been reared from V. edule in Alberta (Lonsdale 2017). The two other known Viburnum miners are both moths that form blotch mines. Phyllonorycter viburnella (Braun) (Gracillariidae), which like M. viburnella is common on V. dentatum on Nantucket Island, forms an underside tentiform mine in which pupation takes place (Braun 1923). Coleophora viburniella Clemens (Coleophoridae) feeds from a portable case, producing full-depth mines that contain no frass. It is recorded from V. nudum L. var. cassinoides (L.) Torr. & A. Gray (McDunnough 1942), V. prunifolium L. (Clemens 1861), and V. rufidulum Raf. (Covell 1999), and we have found larvae on V. dentatum in Vermont. It is conceivable that Marmara mines on Viburnum species other than V. dentatum represent moths distinct from M. viburnella, but at this point we have no reason to suspect so. In Berkshire County, Massachusetts, we have found mines on V. lantanoides in the immediate vicinity of mines on V. dentatum.

Discussion

Among the described Marmara species with known life histories, M. viburnella is unique in its habit of beginning as a leafminer and completing development as a stem miner. Marmara arbutiella is normally a leafminer, occasionally mining in the stem and rarely mining through the stem from one leaf to another (Wagner et al. 2000). Marmara gulosa may mine stems, leaves, or fruits of its various hosts (Guillén et al. 2001). As far as is known, each of the other described species is found in a single plant tissue (stem, leaf, or fruit). However, our observations of leaf mines throughout the USA have revealed that life histories like that of M. viburnella are found on a number of plants unrelated to Viburnum (Table 1). Specifically, confirmed or possible Marmara mines leading from the leaf blade to the stem occur on Anacardiaceae (Malosma Nutt. ex Abrams, Rhus L.), Asteraceae (Lactuca L.), Ericaceae (Arctostaphylos Adans.), Rosaceae (Heteromeles M. Roem.), Salicaceae (Populus L. and Salix L.), and Symplocaceae (Symplocos Jacq.). In Table 1 we have collected records of Marmara mines, or suspected Marmara mines, on North American plants belonging to 115 genera in 50 families. Records from 49 genera in 28 families have been attributed to the polyphagous species M. gulosa, although few of these have been confirmed by rearing or barcoding. Host plant records from 79 genera in 40 families represent the 19 other described North American species plus an unknown number of undescribed species—these records are from outside the known geographic distribution of M. gulosa, or else involve species that have been reared and are known to be undescribed. Apart from M. auratella, which has been reared from two genera of Asteraceae, and M. oregonensis Fitzgerald, which has been reared from two genera of Pinaceae, every other species is monophagous as far as is known (mines on both Castanea Mill. and Quercus L. have been attributed to M. fulgidella, but this species has apparently only been reared from the latter host). Conversely, three described species have been reared from a single host, Fraxinus pennsylvanica Marshall

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 203 (Oleaceae), and two from Malus pumila Mill. (Rosaceae). Several other plant genera host Marmara species that differ sufficiently in geography or larval habit that we suspect multiple moth species are involved. Given all this, it seems likely that undescribed Marmara species far outnumber those that have been named. DRD currently recognizes (and has illustrated) approximately ten new species for North America, most of which have not been reared; the rest of us have reared adults that surely represent more than a dozen further species. It should be noted that mines of other insects can superficially resemble those of Marmara; thus some qualifying statements (e.g. “moth genus not confirmed”) are entered into the “Other notes” field in Table 1. Larvae of the gracillariid genera Phyllocnistis Zeller (Phyllocnistinae) and Metriochroa Busck (Oecophyllembiinae), like Marmara, are sap feeders throughout development. Their leaf mines are similarly long and linear, without granular frass, but in these genera pupation always takes place at the end of the mine, whereas all known leaf-mining Marmara species vacate their mines before spinning cocoons (Kawahara et al. 2017). A few agromyzid flies also produce epidermal linear leaf mines easily mistaken for those of Gracillariidae. These include Phytomyza opacae Kulp, which mines in leaves of evergreen Ilex L. species (Aquifoliaceae) (its puparium is formed within the leaf but is often difficult to detect, as it is formed about 1 cm beyond the apparent end of the mine), and the polyphagous Liriomyza schmidti (Aldrich), which occurs from Florida to Argentina. An unknown agromyzid forms epidermal mines on Gelsemium sempervirens (L.) W.T. Aiton (Gelsemiaceae) in the southeastern USA; we had considered these as possible Marmara mines until one was found with a dead larva inside. The lack of a head capsule easily distinguishes an agromyzid larva from that of a gracillariid, and when a visible frass trail is present, it tends to alternate along the sides of agromyzid mines rather than forming a central line as in gracillariid mines. A wide variety of Lepidoptera, Diptera, Coleoptera, and Hymenoptera feed in stems. Many of these form galleries partially or entirely in the pith, rather than mining exclusively in the outer tissues as in Marmara. We are unaware of any beetles or sawflies that form stem mines resembling those of Marmara. Among flies, some Agromyzidae form externally visible stem mines on herbaceous plants. Most of these are species of Ophiomyia Braschnikov, which pupate at the ends of their mines. Occasional members of other agromyzid genera form stem mines and exit to pupate; as with leaf mines, the frass pattern helps to distinguish these from lepidopteran stem mines. Species of Phytobia Lioy (Agromyzidae) feed in the cambium of woody plants but their mines are not externally visible (Spencer & Steyskal 1986). Apart from Marmara, the moths most likely to be responsible for externally visible stem mines in North America are Nepticulidae. Species of Acalyptris Meyrick form stem mines on Cyperaceae and Fabaceae (Frohne 1939; Wagner 1987), and as far as is known all species of Zimmermannia Hering mine in bark of woody plants, the known hosts being in Fagaceae (we report a possible Zimmermannia mine on Betula L.(Betulaceae) in Table 1, and one European species feeds on Ulmus L. (Ulmaceae)) (van Nieukerken et al. 2016). On Prosopis velutina Wooton (Fabaceae) in Arizona, we have found bark mines resembling those of Z. bosquella (Chambers) on Quercus, each beginning with a compact spiral. None of these nepticulid mines has been reported to exceed 10 cm in length, whereas Marmara stem mines typically measure at least several dm. The other nepticuloid family, Opostegidae, also includes stem miners. The only Nearctic species with a known life history is Opostegoides scioterma (Meyrick), which forms cambium mines on Ribes L. (Grossulariaceae) that are not externally visible (Grossenbacher 1910). Young larvae of the European species of Pseudopostega Kozlov, which are found on Lamiaceae, produce externally visible mines before tunneling deeper into the stems (E. van Nieukerken in litt.). In Massachusetts we have found similar mines on Lycopus L., sometimes extending into the leaf blades, that may likewise be attributable to Pseudopostega. Numerous leaf mines on Monarda L. photographed by C. Vispo (in litt.) in Connecticut are similar in appearance and invariably involve the petioles; these are evidently likewise extensions of stem mines and are likely produced by the same or a related insect. A few moths in other families are known to produce stem mines. Cydia pseudotsugae (Evans) (Tortricidae) forms a mine in the bark of Pseudotsuga menziesii (Mirb.) Franco (Pinaceae) up to 9 dm long, pupating under a semicircular bark flap as in some Marmara but several cm back from the end of the mine (Evans 1969). The European species Leucoptera spartifoliella (Hübner) (Lyonetiidae) has been introduced in western North America, where it forms stem mines on Cytisus Desf. and Genista L. (Fabaceae). The larvae exit their mines to spin conspicuous, elongate white cocoons on the stems (Powell & Opler 2009; Pitkin et al. 2016). Finally, some gracillariids other than Marmara may mine stems. An introduced leaf-mining moth in the southern USA, Caloptilia triadicae Davis, has been observed to mine petioles and stems of Triadica sebifera (L.) Small (Euphorbiaceae) when population densities are high (M. Fox & J. Schneider, in litt.). Powell & Opler (2009)

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A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 207 et al. et al. et al. & *$:>8L!0 ' .* ! . "/5 ( 7887 &, B &/ 4 788: ( 7887 )/, " + " 78:7! . '$ :>G: ( 7887 7888 4 7887!. 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. arbutiella arbutiella M. *$ M. arbutiella arbutiella M. *$ M. gulosa M. +. M. gulosa M. +. gulosa M. +. gulosa M. +. arbutiella M. *$ arbutiella M. *$ M. M. *)24 78:;3 &2)/,+0'*3 )/, ! . Arctostaphylos arbutiella . + . ! ! / ' ))'< . / ) () )' # ))'! ) $ ) & - )' # ) & ( ))' % )'! % $ 78883 et al. % / ) B " ))' )+ ' K )'J/ ) B & ) ! " ))' % ,. )+ 23 2' # 2 ' ! ( $ '! ' % ))'< . + # 2" +' '4 '4 2 2 3 " ))' & Ipomoea purpurea purpurea Ipomoea amomum Cornus Cornus sericea occidentalis 35 4 Nyssa sylvatica Nyssa Dudleya * +& lanatus Citrullus 2" 3 + ( $ Cucurbita $ A 3 Cyclanthus #%'& arizonica Arbutus 3/ 4 menziesii Arbutus unedo Arbutus Arctostaphylos Arctostaphylos andersonii patula Arctostaphylos 4 2) 3 # # # # # ) ) ) ) ) ) ) ) ) , , , , , ,

208 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. et al. & . *$:>:;!. 4 :>>G!( 7887 ( 7887 ( 7887 ( 7887 )/, 0'* " /5 . )/, +0'* +0'* *$:>88 , :>:G , 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. gulosa M. +. M. gulosa M. +. gulosa M. +. gulosa M. +. M. guilandinella *$ M. fulgidella 2) 3 Caloptilia ! ! $ 21'3 ! . $ 2:>M@3 &4 1 Marmara / 78:@ ! )/, * + )2.3 *$ 2:>:;3 1" ) )"!5 ) 1' " ))' *) ! $) () )" '! )"H "H ' 5 ) ) $ $ ))'< 4 - $ ) ) " ))' % )/, * )5 % )9 # " ))' , ) . " ))' # ( % # * ) 23 23 23 23 , ) 2/3 23 F Guilandina Guilandina =/ # )'

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 209 et al. et al. & 5 :>7; ( 7887 ( 7887 ( 7887 ( 7887 " 78:7 +0'* ! . +0'* # . + 5 $ ( 7887 4 788: 4 788: *$:>:; 4 4 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. fulgidella 2) 3 M. gulosa M. +. M. gulosa M. +. gulosa M. +. gulosa M. +. gulosa M. +. gulosa M. +. M. fulgidella 2) 3 3 Q. prinus Q. Marmara Marmara . 4 *$R2:>:;3 O P O $P 2O $P " + )/, )/, . O ' P " ))" ,. )+ / ))' . + )' # )+ / ) B) '< )' 1' # ) & $ % ) =/' ! $ ) $ " ))' " ))' " ))' % ' ) % ))' 3 2 / " % )) " ))' *$ )9 # " ))' Q. montana Q. EO $PF % 5 ) 2 . % 2$+' 3( Quercus alba Quercus kelloggii Quercus ( prinus Quercus N E F Garrya wrightii Garrya virginiana Hamamelis gracilis Deutzia +< Hydrangea 2 3 japonica Itea texana Carya Juglans 3 nobilis Laurus americana Persea Umbellularia californica 2) 3 # # # # # 5 5 5 5 4 4 - 0 0

210 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. et al. et al. et al. et al & :>>G ( 7887 788: ( 7887 ( 7887 4 '$ :>G: 4 788: " 78:: * :>;> ( 7887 ( 7887 . 4 7887!. ( 7887 5 :>7; ( 7887 4 4 4 4 4 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. gulosa M. +. M. gulosa M. +. gulosa M. +. gulosa M. +. gulosa M. +. isortha M. $32 gulosa M. +. M. gulosa M. +. M. gulosa M. +. gulosa M. +. Marmara P P Myrica " + " +" 278:73 O ) ) $ P . )/, )' "H )' ' ) )' " ))' " ))' # )'< 4 ) )"H ! % ) ) % )''< 3 2 % * B " ))' " ))' % 5 9 . 23 ) % )9 # 2 =/( 3 # " I " ))' " ))' 2 3 =/O O " ))' 2$+' 3( 2$+' 3( Umbellularia californica Umbellularia californica granatum Punica Magnolia × soulangeana * Abutilon theophrasti $ barbadense Gossypium hirsutum Gossypium Hibiscus Hibiscus Malva parviflora Melochia pyramidata cacao Theobroma americana Tilia Waltheria indica carica Ficus alba Morus Morus rubra Morella californica 2) 3 Morella cerifera / 2) 3 # # # # #

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 211 & * :>77! 5B :>G; 5B :>G; ( 7887 5B :>G; 5B :>G; ( 7887 .&. :>L; .4 5B :>G? . :>L; .4 5B :>G?! . .&. + # $ :>L; .4 ( 7887 ( 7887 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. fraxinicola M. * M. basidendroca basidendroca M. 5B M. corticola Fitzgerald fraxinicola M. * +. M. gulosa M. oregonensis M. 5B M. fasciella 2) 3 M. gulosa M. +. gulosa M. +. gulosa M. +. oregonensis M. 5B oregonensis M. 5B . + . Marmara Marmara Fraxinus ! ! ()1"! "H2)/,!"/5!.3 1"2.3 1"2.3 1"2.3 ) fasciellaM. fasciellaM. (P (P (P )5 +=/ (P )" ' ) ) ) 2 3 " ))' 2 3 2 %3 " ))' 2 3 " ))' I. % ( *)& % *) *)&)' B. B. * % / ))' . 9 5 23 " ))' & % & ! 2. ) F E 1 Bougainvillea Bougainvillea spectabilis aculeata Pisonia Forsythia 2 3 americana Fraxinus Fraxinus latifolia pennsylvanica Fraxinus pennsylvanica Fraxinus pennsylvanica Fraxinus uhdei Fraxinus 2 B3 fluminense Jasminum 1 europaea Olea balsamea Abies grandis Abies %.. 3 procera Abies monticola Pinus . %.. Pinus strobus menziesii Pseudotsuga 2 35 brasiliensis 2) 3 # # # # # ( ( # # # # # #

212 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. et al & ( 7887 " 78:7 +0'* ! . )/, +0'* 0'# 7887 ( 7887 4 4 ……continued on the next page next on the ……continued $ Marmara M. gulosa M. +. 1 :>:G M. gulosa M. +. P M. elotella M. ! . ! . . . + " + 5B :>G? )/, 4 ! ))'< . ) ) ))' )'< / .)+ / ) )' ) ) ))' / * . ))' )' ( "(! ) + ) "H $ $ $ $ $ $ % ' % # )'< O # % & $ % % . )' % / ))' % . )& % 4 . " ))' " ))' ' ' ' 2' 23 $+ ) 4 4 Tsuga canadensis Tsuga ) EO $PF heterophylla Tsuga 2& 3/ parryi Penstemon 3'4 4 scandens Plumbago Portulaca oleracea scandens Berchemia 2399 Ceanothus ?arboreus Ceanothus 4 fendleri Ceanothus greggii Ceanothus greggii Ceanothus Ceanothus ?integerrimus ' leucodermis Ceanothus 4 sanguineus Ceanothus # sorediatus Ceanothus $+' spinosus Ceanothus ( 4 4 4 2) 3 # # # # # # # # # # & & & & & & & & & & &

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 213 et al. et al. et al et al & . . 4 7887!. *$:>:? ( 7887 .&. + # $ +0'* 788: 1 :>:G .&. 4 7887!. 788: ( 7887 4 4 ……continued on the next page next on the ……continued $ Marmara M. pomonella pomonella M. *$ M. elotella 2*$3 gulosa M. +. M. gulosa M. +. % @I>P ! ))'2.3 2.3! $ $! . + . ' )+ ))' ' ) ) ) ),. )+ ' ))' * )& / - ) ) / )+ / ) )' )5 )+ # ))' "H 2 3 % % $ $ % 2 3 % % % I. 5 9 % $ ' ) % )' 5 9 % . ) ) R )' . )' 4 ! ' " ))' )"2.3 )/, 4 23 23 2 3 " ))' O P * Ceanothus thyrsiflorus thyrsiflorus Ceanothus , arborescens Colubrina 2 3/ asiatica Colubrina californica Frangula 2,3'4 purshiana Frangula 2.)3'4 lupuloides Gouania = obtusifolia Ziziphus 2$%" +' 3'4 4 montanus Cercocarpus arbutifolia Heteromeles 2 3 & arbutifolia Heteromeles 2 3 & arbutifolia Heteromeles 2 3 & Malus pumila Malus EO PF pumila Malus EO PF Photinia 2 3 & 2) 3 # # # # # & & & & & & & & & & & & & & &

214 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. et et al. :>>8!" & ( 7887 ( 7887 ( 7887 ( 7887 ( 7887 ( 7887 *$:>:? al +" 78:7 !. .* 788: ( 7887 0'# 4 4 4 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. gulosa M. +. gulosa M. +. gulosa M. +. gulosa M. +. gulosa M. +. M. serotinella *$ gulosa M. +. M. gulosa M. +. )" ' - ()2)/,!"/5!.3 . ! . . + )"H . 5 ) 1' )" +# ) )' )+ / ) )' / ))' $ $ % " ))' % " ))' " ))' )' " ))' ) " ))' % ) ) ) 4 % ' ) " ))' ) 2( , ! 23 23 ) + 2 3 nucipersica persica Rubus Rubus F % ' )+ 2 3 " ))' Prunus Prunus 2 3 caroliniana Prunus ' dulcis Prunus .'EO PF ilicifolia Prunus %$+' 3. . laurocerasus Prunus pensylvanica Prunus persica Prunus * 2/$3)9/ EO PF persica Prunus * EO PF serotina Prunus Rosa Rosa californica +/ Rubus armeniacus Rubus 5$E procerus Rubus ursinus / 2) 3 # # # # # & & & & & & & & & & & & & &

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 215 et al. et al. & '$ :>@:! 4 788: '$ :>G: '$ :>G: ( 7887E / 78:8F '$ :>G:! ( 7887E / 78:8F 4 :>>G ( 7887E / 78:8F . . )" . . .&. + # $ 4 4 4 4 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. gulosa M. +. gulosa M. +. +. M. gulosa M. +. +. +. M. gulosa M. gulosa M. +. gulosa M. gulosa M. 2 2 3 ! ! ! ! )' )' ) ) ))' ) ) "H 5 )' 1 )' 1 =/ % & ) & ) )' % ' )+ % 5! 9 % (( 9 / 0 A / 0 A % # )'< *% )"H . + F F 23 % 5 9 * 0 23 23 ) 2* 23 ) medica × aurantifolia sinensis × maxima 2 3 & Rubus vitifolius vitifolius Rubus +/ alba Chiococca parvifolia Chiococca %4 fruticosa Erithalis ×aurantiifolia Citrus 2) 3/ 2 3E &F 2 ×limon Citrus * 2 3 E Citrus maxima 3 2 3 ×paradisi Citrus 2 3 E 2 3 reticulata Citrus 2 3 ×sinensis Citrus $2 3 E % & F 21 3 ×tangelo Citrus - +, 2 3 trifoliata Poncirus 2) 3 # # # # # & & & & & & & & & & & &

216 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. et al. et al. & . ( 7887 ) :M@; . 788: ( 7887 .&. )/, +0'* 78:? . 4 )/, +0'* 788: " + " 78:: ! . 4 4 4 ……continued on the next page next on the ……continued $ Marmara M. salictella ) M. gulosa M. +. M. salictella ) O M. salictella ) gulosa M. +. M. gulosa M. +. M. salictella ) O ! M. gulosa Salix . . + . 4 ! % . ! "/5 2 3 2 ! ) # )+ / ))' )"H )5 / ))' - 1" & / ) () )"H ))' / ) )' % ) % ) )+ % % ) ) ) ) 0 ) % =/ . )+ )' 4 23 " " ))' * * % ) ) )+ ( $ ' % * .! ) )+ 2 3 EO PF % # )+ " ))' '&- 1 :>:G!. EO ! / Zanthoxylum fagara fagara Zanthoxylum Populus Populus Populus grandidentata Populus % tremuloides Populus % trichocarpa Populus " +'4 % $ Salix Salix PF Salix lasiolepis Salix lasiolepis nigra Salix purpurea Salix floridanum Acer 2) 3# % macrophyllum Acer # negundo Acer palmatum Acer 2) 3 # # # # # & / / / / / / / / / / / / / / / /

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 217 et al. & ) :MG? # .&. + .$ ( 7887 ( 7887 )/, ! "/5 ! . . .&. + # $ ( :>7M!"/ 5 ) 78:L!)/ , +0' !"/ * 5 ! 78:@ "/5 ! # "/5 "/5 ! . 4 4 ……continued on the next page page next on the ……continued $ Marmara M. gulosa M. +. gulosa M. +. M. smilacisella 2) 3 M. smilacisella 2) 3 M. smilacisella 2) 3 M. smilacisella 2) 3 M. smilacisella 2) 3 M. smilacisella 2) 3 % B glabra Smilax &% ' . ( ' !$ )" ! ( ' 2)/,3 9 2 =/( 3 )"! '! / ) () )5 () ' )5! # )-(! * .! $ ) ) ()!# 9 / ) () ()! )"H # )9 ! 9P % 5 9 " ))' % / ) .) % I. % / ) )+ 2$ / ) % ) " ))' ) )'! drummondii drummondii saponaria Acer pseudoplatanus pseudoplatanus Acer rubrum Acer rubrum Acer saponaria Sapindus saponaria Sapindus glauca Simarouba Smilax bona-nox Smilax glabra Smilax glauca Smilax Smilax laurifolia Smilax rotundifolia +' 3.* 2) 3 # # # # # / / / / / / / / / / / /

218 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. et al. et al. & / :>G:! . ( 7887 78:8 ( 7887 * :>8> ( 7887 788:!#' ( 7887 "/5 "/5 5 :>7; 4 :>>G 4 4 4 4 4 4 $ Marmara +. +. +. M. smilacisella 2) 3 M. smilacisella 2) 3 gulosa M. gulosa M. gulosa M. +. gulosa M. M. gulosa M. +. gulosa M. +. M. elotella 5B + 4 . K )/, ! ) )"! (P! ) ) 1"!* B )"H () 2 ' )3 )' ()!* $ )/) $ % ) )' % ) )5 / ) % " ))' !O P " ))' . # )'< " ))' " ))' 23 . " ))' F ! '4 '4 % ' ) / ) E Q& F EO PF O 1' 2 S. hispida S. . Smilax smallii smallii Smilax tamnoides Smilax E annuum Capsicum Datura inoxia Datura meteloides glauca Nicotiana 4 erianthum Solanum melongena Solanum xanti Solanum tinctoria Symplocos S Ulmus americana Ulmus vinifera Vitis 4R vinifera Vitis Q" /R Q5 R Vitis 3 2) 3 # # # # # / / / / / / / / / = = 1 1 1

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 219 report a coastal California Phyllocnistis species on Salix L. (Salicaceae) whose mines sometimes extend from leaf to leaf along the stems. In Europe, the Salix feeder P. ram ul icola Langmaid & Corley forms a very Marmara-like stem mine, up to 30 cm long, only entering a leaf blade to spin its cocoon, which is done along the leaf margin adjacent to the petiole (Langmaid & Corley 2007). In light of these examples, empty stem mines can only confidently be attributed to Marmara when they terminate in the characteristic bark flap or if larval exuviae can be removed and examined from the mine—the head capsule of Marmara is sufficiently diagnostic as to allow certain identification.

Acknowledgments

We thank the Nantucket Biodiversity Initiative for financial and other support for our ongoing leafminer surveys on Nantucket; the Nantucket Conservation Foundation and Massachusetts Department of Conservation for permission to collect specimens and dig up viburnums on their properties; Erik van Nieukerken (Naturalis Biodiversity Center, Leiden, Netherlands) for sharing his observations of stem-mining Nepticuloidea; Robert L. Zuparko (Department of Entomology, California Academy of Sciences, San Francisco, United States) for identifying the encyrtids; Christer Hansson (Department of Zoology, Lund University, Sweden) for identifying the eulophids; Blaine Mathison (Salt Lake City, Utah, United States) and Vassili Belov (Department of Entomology, Texas A&M University, United States) for identifying photographed beetles; Frans Janssens (Antwerp, Belgium) for identifying photographed springtails; Karolyn Darrow for the photograph of the holotype and Young Sohn for the illustration of the male genitalia (Department of Entomology, Smithsonian Institution, United States); Shigeki Kobayashi (Osaka Prefecture University, Japan) and an anonymous reviewer for feedback that improved the manuscript.

References

Atkins, E.L. (1961) Citrus peel miner. California Citrograph, 46, 367–371. Atkins, E.L. (1971) The citrus peel miner, more problems in the desert. California Citrograph, 56, 245–246 & 267. Bondar, G. (1939) Insetos daninhos e parasitas do cacau. Livraria duas Americas, Bahia, Series pragas e molestias, Boletim Tecnico, 5, 69–73. Braun, A.F. (1909) Notes on Chambers’ of Tineina. Entomological News, 20, 428–434. Braun, A.F. (1915) New genera and species of Tineina. The Canadian Entomologist, 47 (6), 188–197. https://doi.org/10.4039/Ent47188-6 Braun, A.F. (1922) Microlepidoptera: notes and new species. The Canadian Entomologist, 54, 90–94. [species] https://doi.org/10.4039/Ent5490-4 Braun, A.F. (1923) Microlepidoptera: notes and new species. Transactions of the American Entomological Society, 49 (2), 115– 127. Busck, A. (1900) New species of moths of the superfamily Tineina from Florida. Proceedings of the United States National Museum, 23, 225–254. https://doi.org/10.5479/si.00963801.23-1208.225 Busck, A. (1904) Tineid moths from British Columbia, with descriptions of new species. Proceedings of the United States National Museum, 27, 745–778. https://doi.org/10.5479/si.00963801.27-1375.745 Busck, A. (1907) New American Tineina. Proceedings of the Entomological Society of Washington, 8 (3–4), 86–99. Busck, A. (1909) Notes on microlepidoptera, with descriptions of new North American species. Proceedings of the Entomological Society of Washington, 11 (2), 87–103. Busck, A. (1913) Note on a barkmining lepidopteron of the genus Marmara Clemens. Proceedings of the Entomological Society of Washington, 15 (4), 150. Busck, A. (1915) Descriptions of new North American microlepidoptera. Proceedings of the Entomological Society of Washington, 17 (2), 79–94. Carr, J.F. (2014) Vine mine. Available from: http://bugguide.net/node/view/910263 (accessed 7 July 2017) Chambers, V.T. (1875) Tineina of the central United States. The Cincinnati Quarterly Journal of Science, 2 (2), 97–121. Clemens, B. (1861) Micro-lepidopterous larvae. Notes on a few species, the imagos of which are probably undescribed. Proceedings of the Entomological Society of Philadelphia, 1 (4), 75–87. Clemens, B. (1863) American micro-lepidoptera. Proceedings of the Entomological Society of Philadelphia, 2 (1), 4–14. Covell, C. (1999) The butterflies and moths (Lepidoptera) of Kentucky: an annotated checklist. Kentucky State Nature Preserves Commission Scientific and Technical Series, 6, 1–220.

220 · Zootaxa 4337 (2) © 2017 Magnolia Press EISEMAN ET AL. Davis, D.R., McKay, F., Oleiro, M., Vitorino, M.D. & Wheeler, G.S. (2011) Biology and systematics of the leafmining Gracillariidae of Brazilian pepper tree, Schinus terebinthifolius Raddi, with descriptions of a new genus and four new species. Journal of the Lepidopterists’ Society, 65 (2), 61–93. https://doi.org/10.18473/lepi.v65i2.a1 De Benedictis, J.A., Wagner, D.L. & Whitfield, J.B. (1990) Larval hosts of microlepidoptera of the San Bruno Mountains, California. Atala, 16 (1–2), 14–35. De Gryse, J.J. (1943) Note on Marmara fasciella Chambers. The Canadian Entomologist, 75 (2), 40. https://doi.org/10.4039/Ent7540-2 Ely, C.R. (1917) A revision of the North American Gracilariidae from the standpoint of venation. Proceedings of the Entomological Society of Washington, 19 (1–4), 29–77. Evans, D. (1969) Laspeyresia pseudotsugae n. sp. (Lepidoptera: Olethreutidae), a bark miner in Douglas-fir. The Canadian Entomologist, 101, 955–963. https://doi.org/10.4039/ent101955-9 Fitzgerald, T.D. (1973) Coexistence of three species of bark-mining Marmara (Lepidoptera: Gracillariidae) on green ash and descriptions of new species. Annals of the Entomological Society of America, 66 (2), 457–464. https://doi.org/10.1093/aesa/66.2.457 Fitzgerald, T.D. (1975) A new species of bark-mining Marmara (Lepidoptera: Gracillariidae) from Douglas-fir. Annals of the Entomological Society of America, 68 (3), 545–548. https://doi.org/10.1093/aesa/68.3.545 Fitzgerald, T.D. & Simeone, J.B. (1971) Serpentine miner Marmara fraxinicola (Lepidoptera: Gracillariidae) in stems of white ash. Annals of the Entomological Society of America, 64 (4), 770–773. https://doi.org/10.1093/aesa/64.4.770 Forbes, W.T.M. (1923) The Lepidoptera of New York and neighboring states. Primitive forms, microlepidoptera, pyraloids, bombyces. Cornell University Agricultural Experiment Station Memoir, 68, 1–729. https://doi.org/10.5962/bhl.title.23875 Frohne, W.C. (1939) Observations on the biology of three semiaquatic lacustrine moths. Transactions of the American Microscopical Society, 58 (3), 327–348. https://doi.org/10.2307/3222884 Gates, M.W., Heraty, J.M., Schauff, M.E., Wagner, D.L., Whitfield, J.B. & Wahl, D.B. (2002) Survey of the parasitic Hymenoptera on leafminers in California. Journal of Hymenoptera Research, 11 (2), 213–270. Gibson, R., Bacon, D., Langston, D., Kerns, D. & Gibson, R. (1997) The Citrus Peel Miner, Marmara salictella, in Arizona Grapefruit in 1994. 1997 Citrus Research Report, University of Arizona, College of Agriculture Series P-109, 1–6. Gilmore, S. (2013) Leaf Miner Moth 2. Available from: http://bugguide.net/node/view/773621 (accessed 7 July 2017) Grossenbacher, J.G. (1910) Medullary spots: a contribution to the life history of some cambium miners. New York Agricultural Experiment Station Technical Bulletin, 15, 49–65. Guillén, M., Davis, D.R. & Heraty, J.M. (2001) Systematics and biology of a new, polyphagous species of Marmara (Lepidoptera: Gracillariidae) infesting grapefruit in the southwestern United States. Proceedings of the Entomological Society of Washington, 103 (3), 636–654. Haines, D. (2010) Marmara gulosa Guillen & Davis. Available from: http://bugguide.net/node/view/374351 (accessed 21 December 2016) Kawahara, A.Y., Plotkin, D., Ohshima, I., Lopez-Vaamonde, C., Houlihan, P.R., Breinholt, J.W., Kawakita, A., Xiao, L., Regier, J.C., Davis, D.R., Kumata, T., Sohn., J.-C., de Prins, J. & Mitter, C. (2017) A molecular phylogeny and revised higher-level classification for the leaf-mining moth family Gracillariidae and its implications for larval host-use evolution. Systematic Entomology, 42, 60–81. https://doi.org/10.1111/syen.12210 Langmaid, J.R. & Corley, M.F.V. (2007) Phyllocnistis ramulicola sp. nov. (Lepidoptera: Gracillariidae) in England and Portugal. Entomologist’s Gazette, 58, 227–237. Lonsdale, O. (2017) The Liriomyza (Agromyzidae: Schizophora: Diptera) of Canada & Alaska. Zootaxa, 4234 (1), 1–156. https://doi.org/10.11646/zootaxa.4234.1.1 Mallory, C. (2013) ?Marmara mine Laurel Sumac. Available from: http://bugguide.net/node/view/869516 (accessed 21 December 2016) McDunnough, J. (1942) Further notes on maritime Coleophoridae (Lepidoptera). The Canadian Entomologist, 74, 167–172. https://doi.org/10.4039/ent74167-9 Needham, J.G., Frost, S.W. & Tothill, B.H. (1928) Leaf-mining Insects. The Williams & Wilkins Company, Baltimore, Maryland, 351 pp. Neff, D.C. (2002) Ecological studies of citrus peelminer, Marmara gulosa Guillén and Davis, in the San Joaquin Valley. M.S. thesis, California Polytechnic State University, San Luis Obispo, 78 pp. Nieukerken, E.J. van, Doorenweerd, C., Hoare, R.J.B. & Davis, D.R. (2016) Revised classification and catalogue of global Nepticulidae and Opostegidae (Lepidoptera, Nepticuloidea). ZooKeys, 628, 65–246. https://doi.org/10.3897/zookeys.628.9799 Pitkin, B., Ellis, W., Plant, C. & Edmunds, R. (2016) The leaf and stem mines of British flies and other insects. Available from:

A NEW SPECIES OF MARMARA Zootaxa 4337 (2) © 2017 Magnolia Press · 221 http://www.ukflymines.co.uk/index.php (accessed 4 April 2017) Powell, J.A. (2012) Lepidoptera recorded on Santa Catalina Island. Available from: https://essig.berkeley.edu/documents/ lepidoptera/santa_catalina_island_jerry_powell.pdf (accessed 21 December 2016) Powell, J.A. & Opler, P.A. (2009) Moths of Western North America. University of California Press, Berkeley, 383 pp. Semet, P. (2010) Taxonomy, biogeography and pest status of Marmara sp. in northern Mexico and California citrus production. M.S. thesis, California Polytechnic State University, San Luis Obispo, 61 pp. Simon, A. (2016) Salal (Gaultheria shallon) leaf miner? Available from: http://bugguide.net/node/view/1278156 (accessed 21 December 2016) Spencer, K.A. & Steyskal, G.C. (1986) Manual of the Agromyzidae (Diptera) of the United States. U.S.D.A. Agriculture Handbook, 638, 478 pp. Stevens, P.F. (2016) Angiosperm Phylogeny Website. Available from: http://www.mobot.org/MOBOT/research/APweb/ (accessed 21 December 2016) Turner, L. & Turner, J. (2011a) Marmara salictella. Available from: http://bugguide.net/node/view/603638 (accessed 21 December 2016) Turner, L. & Turner, J. (2011b) Marmara sp. Available from: http://bugguide.net/node/view/603978 (accessed 21 December 2016) Turner, L. & Turner, J. (2012a) Marmara sp. Available from: http://bugguide.net/node/view/607409 (accessed 21 December 2016) Turner, L. & Turner, J. (2012b) Marmara sp. Available from: http://bugguide.net/node/view/607535 (accessed 21 December 2016) Turner, L. & Turner, J. (2012c) Marmara sp. Available from: http://bugguide.net/node/view/607548 (accessed 21 December 2016) USDA, NRCS (2016) The PLANTS Database. National Plant Data Team, Greensboro, NC. Available from: http:// plants.usda.gov (accessed 21 December 2016) Vinal, S.C. (1917) Notes on the life-history of Marmara elotella Busck, a lepidopterous sap feeder in apple twigs. Journal of Economic Entomology, 10, 488–496. https://doi.org/10.1093/jee/10.5.488 Wagner, D.L. (1987) A new Microcalyptris species from California (Lepidoptera: Nepticulidae). Pan-Pacific Entomologist 63 (3), 278–283. Wagner, D.L., Loose, J.L., Fitzgerald, T.D., DeBenedictis, J.A. & Davis, D.R. (2000) A hidden past: the hypermetamorphic development of Marmara arbutiella (Lepidoptera: Gracillariidae). Annals of the Entomological Society of America, 93 (1), 59–64. https://doi.org/10.1603/0013-8746(2000)093[0059:AHPTHD]2.0.CO;2 Wilson, T. (2014) Viburnum Leaf Mine ID Request. Available from: http://bugguide.net/node/view/1014804 (accessed 21 December 2016) Wilson, T. (2015) Boxelder Stem Miner(?) ID Request. Available from: http://bugguide.net/node/view/1067950 (accessed 21 December 2016) Wilson, T. (2016) Greenbrier Leaf Miner [=Marmara smilacisella?] ID Request. Available from: http://bugguide.net/node/ view/1307163 (accessed 7 July 2017)